PLANT-WATER
RELATIONS
Plant Cells & Water
Water Transport Processes
There are two major ways to move H2O
molecules:
Diffusion
Bulk (or Mass) Flow
Diffusion
net, random movement of individual molecules from
one area to another.
molecules move from high to low concentration,
following a concentration gradient.
Thermodynamically, the molecules move from an area
of high free energy (i.e., higher concentration) to one of
low free energy (i.e., lower concentration).
The net movement stops when a dynamic equilibrium
is achieved.
Bulk (or Mass) Flow
the concerted movement of groups of molecules en
masse, in response to a pressure gradient.
molecules move from high to low pressure, following a
pressure gradient.
Examples:
1. faucet -- When you turn a faucet on, water comes
out. This occurs because the water in the tap is under
pressure relative to the air outside the faucet.
2. toilet -- high pressure in the tank/bowl but lower
pressure in the sewer system.
Osmosis
specialized case of diffusion
represents the diffusion of a solvent (typically water)
across a membrane.
Dialysis
another specialized case of diffusion
the diffusion of solute across a semi-permeable
membrane.
Example – consider a cell containing a sugar dissolved in
water. If water (the solvent) moves out of the cell into
the surroundings it moves osmotically; if the sugar
(solute) moves into the surroundings, it is an example
of dialysis.
The movement of water across a membrane is
a combination of diffusion and bulk flow.
Individual water molecules diffuse across the
membrane.
There are integral proteins (called aquaporins) in the
membrane that form a channel or pore through which
water moves. Water molecules move through these
pores by bulk flow. Essentially, the protein pores serve
as water transport channels.
Water flow is a passive process, in response to physical
forces and following a gradient of free energy.
Factors influencing the rate of
diffusion
Several factors influence the rate of diffusion. These include:
Concentration Gradient
Molecular Speed
Temperature
Pressure
Solute effect on the chemical potential
of the solvent
A. Concentration Gradient
the rate of diffusion is directly proportional to the
concentration gradient.
The greater the difference in concentration between
two areas, the greater the rate of diffusion.
Thus, when the gradient is zero, there will be no net
diffusion. Diffusion will only occur so long as a
concentration gradient exists.
B. Molecular Speed
According to kinetic theory, particles like atoms
and molecules are in always in motion at
temperatures above absolute zero (0 K = -
2730C).
Molecular movement is:
directly proportional to temperature.
related to MW (heavier particles move more slowly
than lighter, smaller ones).
C. Temperature
increases the rate of molecular movement, therefore,
increases the rate of diffusion.
D. Pressure
increases speed of molecules, therefore, it increases the
rate of diffusion.
E. Solute effect on the chemical potential of
the solvent (water)
Solute particles decrease the free energy of a solvent.
critical factor: number of particles, not the charge or
particle size.
mole fraction of solvent = # solvent molecules/
total molecules (# solvent molecules + # solute
molecules).
Essentially, solvent molecules, such as water in a
biological system, move from a region of greater mole
fraction to a region where it has a lower mole fraction.
Water diffuses from an area of higher mole fraction or
higher energy to an area of lower mole fraction or
lower energy.
Water Potential
measure of the energy state of water.
important concept in plant physiology because
it determines the direction of movement and
status of water in plant cells.
A. Some Definitions
Free energy - energy available to do work.
(thermodynamic property defined as the energy isothermally available for
conversion to work).
Chemical potential (µ) - free energy/unit quantity of a substance
(usually per mole); (J mol-1).
Water potential (Ψw) - chemical potential of water, compared to pure
water at the same temperature and pressure. The units are in pressure
because:
(a) plant cells under pressure (remember the wall?)
(b) it is easier to measure pressure.
Pressure is measured in MPa (megapascals).
1 MPa = 10 bars = 10 atm.
B. Components of water potential
1. Solute (or osmotic) potential (Ψs)
This is the contribution due to dissolved solutes.
Solutes always decrease the free energy of water, thus
their contribution is always negative.
The solute potential of a solution can be calculated with
the van’t Hoff equation:
Ψs = - miRT
where: m = molality (moles/1000 g); i = ionization
constant (often 1.0); R = gas constant (0.083 liter x
bars/mol deg); and T = temperature (K).
2. Pressure (or Pressure Potential, Ψp)
Due to the pressure build up in cells (turgor).
(thanks to the cell wall).
It is usually positive, although may be negative (tension)
as in the xylem.
3. Matric potential (Ψm)
This is the contribution due to the force of attraction of
water for colloidal, charged surfaces.
It is negative because it reduces the ability of water to
move.
In large volumes of water, it is very small and usually
ignored.
However, it can be very important in the soil, especially
when referring to the root/soil interface.
4. Gravity (Ψg)
Contributions due to gravity which is usually ignored,
unless referring to the top of tall trees.
Water Enters or Leaves the Cell along a
Water Potential Gradient.
The direction and rate of water flow across a
membrane are determined by the sum of:
concentration gradient of water
pressure gradient
Water Transport Rates Depend on Driving
Force and Hydraulic Conductivity.
The direction of flow is determined by the direction of
the Ψw gradient (the driving force for transport).
The rate of water movement is proportional to:
1. magnitude of Ψw gradient
2. hydraulic conductivity of the membrane
Flow rate = driving force x hydraulic conductivity
Ψw serves 2 functions:
1. to predict patterns of H2O movement
It is the Ψw that drives water movement through
plants.
This is a result of physical forces, no direct energy
input into moving water.
2. to monitor plant H2O status
The Water Potential
Concept Helps Us
Evaluate the Water
Status of a Plant.
Plants can change the Ψw of the cell
there is an upper limit ~-0.5 MPa;
typically Ψs = -0.8 to -1.2 MPa
Plants can increase osmotic concentration.
(make Ψs more negative) in order to lower Ψw to allow
the plant to extract water.
-- in response to drought and salinity stress.
-- plants often synthesize higher levels of proline as a
stable osmoticum, and other compatible solutes.
Ψw is affected by all factors which change the
free energy or chemical activity of water
molecules.
Ψw is increased (made less negative) by:
1. dev’t of turgor pressure
2. increase in temperature
Ψw is decreased (made more negative) by:
1. addition of solutes
2. matric forces which absorb or bind water
3. negative pressure (tension)
4. reduction in temperature
In the war of sun and dryness against living things, life has its
secrets of survival. Life no matter on what level, must be
moist or it will disappear.
–John Steinbach
